CIHM 
Microfiche 
Series 
(Monographs) 


ICMH 

Collection  de 
microfiches 
(monographies) 


Canadian  l.irtituta  for  Historical  Microraproductions  /  Institut  Canadian  da  microraproductions  historiquas 


Technical  and  Bibliographic  Notes  /  Notes  technique  et  bibliographiques 


The  Institute  has  attempted  to  obtain  the  best  original 
copy  available  for  filming.  Features  of  this  copy  which 
may  be  bibliographically  unique,  which  may  alter  any  of 
the  images  in  the  reproduction,  or  which  may 
significantly  change  the  usual  method  of  filming  are 
checked  below. 


^ 


Coloured  covers  / 
Couvetture  de  couleur 


I     j  Covers  damaged  / 

' — '  Couverture  endonimagee 

I     I  Covers  restored  and/or  laminated  / 

' — '  Couverture  restaurde  et/ou  pellicula 

I     I  Cover  title  missing  /  Le  titre  de  couverture  manque 

I     I  Coloured  maps  /  Cartes  geographiques  en  couleur 

r~j  Coloured  ink  (i.e.  other  than  blue  or  black)  / 

Encre  de  couleur  (i.e.  autre  que  bleue  ou  noire) 

I     I  Coloured  plates  and/or  illustrations  / 

—  Planches  et/ou  illustrations  en  couleur 

I     I  Bound  with  other  material  / 

' — '  Reli6  avec  d'autres  documents 


D 
D 


D 


Only  edition  available  / 
Seule  ^ition  disponible 

Tight  binding  may  cause  shadows  or  distortion 
along  interior  margin  /  La  reliure  serree  peut 
causer  de  I'ombre  ou  de  la  distorsion  le  long  de 
la  marge  int^rieure. 

Blank  leaves  added  during  restoratkxis  may  appear 
within  the  text.  Whenever  possible,  these  have 
been  omitted  from  filming  /  II  se  peut  que  ceitaines 
pages  blanches  ajout^es  lors  d'une  restauration 
apparaissent  dans  le  texte,  mais,  k>rsque  cela  dtait 
possible,  ces  pages  n'ont  pas  ^  filmdes. 


L'Institut  a  microfilme  le  meilleur  examplaire  qu'il  lui  a 
ete  possible  de  se  procurer.  Les  d4tails  de  cet  exem- 
plaire  qui  sont  peut-etre  uniques  du  point  de  vue  bibli- 
ographique,  qui  peuvent  modifier  une  image  reproduite, 
ou  qui  peuvent  exiger  une  modifications  dans  la  m6th- 
ode  normale  de  filmage  sont  indiqu6s  ci-dessous. 

I      I      Coloured  pages  /  Pages  de  couleur 

I      I      Pages  damaged  /  Pages  endommagees 

I      I      Pages  restored  and/or  laminated  / 
' — '      Pages  restaur6es  et/ou  pellk:ul6es 

r^    Pages  discoloured,  stained  or  foxed  / 
' — '      Pages  decolor6es,  tachet6es  ou  piquees 

I     I      Pages  detached  /  Pages  d6tachees 

r^^   Showthrough  /  Transparence 

I     I      Quality  of  print  varies  / 

' — '      Quafite  inegale  de  I'impression 

j     j      Includes  supplementary  material  / 
—      Comprend  du  materiel  :>uppl^mentaire 

I  I  Pages  wholly  or  partially  obscured  by  errata 
' — '  slips,  tissues,  etc.,  have  been  refilmed  to 
ensure  the  best  possible  image  /  Les  pages 
totalement  ou  partiellement  obscurcies  par  un 
feuillet  d'errata,  une  pelure,  etc.,  ont  6\^  filmees 
k  nouveau  de  fagon  a  obtenir  la  meiileure 
image  possible. 

I  I  Opposing  pages  with  varying  colouration  or 
' — '  discolourations  are  filmed  twice  to  ensure  the 
best  possible  image  /  Les  pages  s'opposant 
ayant  des  colorations  variables  ou  des  d^ol- 
orations  sont  filmev^s  deux  fois  afin  d'obtenir  la 
meilleur  image  possible. 


\2f 


Additk>nal  comments  / 
Commentaires  supplementaJres: 


Pagination  is  as  follows  :  p.  81-106. 


This  ittm  it  f  ilmad  at  the  reduction  ratio  chacfcid  balow/ 

Ce  document  est  filmi  au  taux  de  rMuction  indique  ci-detsous. 

10X  14X  18X 


a 


22X 


12X 


16X 


20X 


26X 


30X 


J 


24X 


?8X 


32X 


The  copy  filmed  h«r«  has  baan  raproducad  thanka 
to  tha  ganaroaity  of: 

Harriet  Irving  Litrary 
University  of  New  Brunswick 

The  images  appearing  here  are  the  best  quality 
possible  considering  the  condition  and  legibility 
of  the  original  copy  and  in  keeping  with  the 
filming  contract  specifications. 


Original  copies  in  printed  paper  covers  are  filmed 
beginning  with  the  front  cover  and  ending  on 
the  last  page  with  a  printed  or  illustrated  impres- 
sion, or  tha  back  cover  when  appropriate.  All 
other  original  copies  are  filmed  beginning  on  the 
first  page  with  a  printed  or  illustrated  impres- 
sion, and  ending  on  the  last  page  with  a  printed 
or  illustrated  impression. 


The  last  recorded  frame  on  each  microfiche 
shall  contain  the  symbol  -^  (meaning  "CON- 
TIIMUED").  or  the  symbol  V  (meaning  "END"), 
whichever  applies. 

Maps,  plates,  charts,  etc.,  may  be  filmed  at 
different  reduction  ratios.  Those  too  large  to  be 
entirely  included  in  one  exposure  are  filmed 
beginning  in  the  upper  left  hand  corner,  left  to 
right  and  top  to  bottom,  as  many  frames  as 
required.  The  following  diagrams  illustrate  the 
method: 


L'exemplaira  film*  fut  raproduit  grica  A  la 
g*n«roslt*  da: 

Harriet  Inring  Library 
University  of  New  Brunswick 

Les  images  suivantes  ont  «t«  reproduites  avac  la 
plus  grand  soin,  compta  tenu  de  la  condition  at 
de  la  nettet«  de  l'exemplaira  film*,  et  en 
conformity  avec  les  conditions  du  contrat  de 
filmage. 

Lea  exemplaires  originaux  dont  la  couvarture  en 
papier  est  imprim«e  sont  film«s  en  commenpant 
par  le  premier  plat  et  en  terminant  soit  par  la 
darniire  page  qui  comporte  une  empreinte 
d'impression  ou  d'illustration,  soit  par  le  second 
plat,  salon  le  cas.  Tous  les  autres  exemplaires 
originaux  sont  filmte  en  commenpant  par  la 
premiere  page  qui  comporte  une  empreinte 
d'impression  ou  d'illustration  at  en  terminant  par 
la  dernidre  page  qui  comporte  une  telle 
empreinte. 

Un  des  symboles  suivants  apparaftra  sur  la 
dernidre  image  de  cheque  microfiche,  selon  le 
cas:  le  symbole  — •►  signifie  "A  SUIVRE",  le 
symbols  V  signifie  "FIN". 

Les  cartes,  planches,  tableaux,  etc.,  peuvent  Atre 
filmte  d  des  taux  de  reduction  diffirents. 
Lorsque  le  document  est  trop  grand  pour  Atre 
reproduit  en  un  seul  clich«,  11  est  film«  A  partir 
de  I'angle  supArieur  gauche,  de  gauche  A  droite. 
et  de  haut  en  bas,  en  prenant  le  nombre 
d'images  ndcessai'e.  Les  diagrammas  suivanta 
illustrent  la  m^thode. 


1  2  3 


1  2  3 

4  5  6 

. 


MiaoCOnr   RiSOlUTION   TKT  CHART 

,    NS!  and  ISO  TEST  CHART  No   2) 


^  APPLIED  IM/IGE    Inc 

JBC.  '655  €03l  Main  Streei 

B^S  Roc^ieater.    Naw   Yorl<         l*609        uSA 

^^S  (^'6)  *82  -  0300  -  Phor-e 

^^S  (716)   2S8  -  5989  -  Fa« 


Reprinted  iron   Th*  Bthmital  GattUf,  4t;  8i-io«,  AnguX,  1906 


THE    NASCENT   FOREST  OF   THE   MIS- 
COU  BEACH  PLAIN 

CONTRIBUTIONS  TO  THE  ECOLOGICAL  PLANT  GEOORAPHV  OK  THE 
PROVINCE  OF  NEW  BRUNSWICK.    NO.  4 

(WITH  FO'JKTBBN  nOURES) 


W.  F.  GANON<; 


PSUrrU)  AT  m  UMIVBRUTV  Or  CHICAGO  PKKSS 


VOLUME  XIJI 


NUMIIKK  2 


! 


i 


Botanical   Gazette 


AUGUST,   igo6 

THE  NASCENT  FOREST  OF  THE  MISCOU  BEACH  PLAIN. 

CONTRIBUTIONS   TO   TIIK    KCOI.OOICAL    PLANT    (JEOORAI'HY 
OF  THK  I'ROVINCK  OF  NKVV  HRUNSWICK,  NO.  4.' 
\V.  F.  Canon  o. 

(with  fourtkkn  figures) 

The  extreme  northeastern  angli  of  the  Province  of  New  Hruns- 
wick,  as  the  accompanying  map  will  show,  is  formed  by  the  island 
of  Miscou.  The  northwestern  margin  of  this  island  is  an  extensive 
sandy  beach  plain,  growing  rapidly  by  action  of  the  sea,  locally 
called  Grande  Plaine.  On  this  plain  there  is  developing  a  forest 
which  exhibits  every  stage  of  formation  from  the  salt  plants  of  the 
open  sea  beach  to  the  heterogeneous  vegetation  of  the  mixed  woods. 
The  conditions  are  unusual  and  the  phenomena  of  proportional 
interest.  In  August  1905  I  wa.s  able  to  give  the  place  some  two 
weeks  of  oteervational  study,  with  results  which  follow. 

In  all  such  studies  as  this  the  correct  identification  of  the  plants 
is  of  first  importance,  and  idcntificalun  is  becoming  a  matter  of 
such  difficulty  that  only  a  professional  systematist  is  competent 
authority.  Accordingly  I  have  sent  all  of  my  a  llections,  including 
a  .specimen  of  every  plant  I  found  at  Grande  Plaine,  to  Professor 
M.  L.  Fernald,  of  the  Gray  Herbarium  of  Harvard  University, 
who  has  been  so  kind  as  to  determine  their  identity,  and,  as  well, 
to  give  me  the  names  they  should  bear  in  accordance  with  the  recom- 
mendations of  the  Vienna  Congross.  I  wish  here  to  express  my 
indebtedness  to  him  and  my  best  thanks  for  this  invaluable  aid. 
Such  is  the  origin  of  the  nomenclature  of  this  paper. 

■  No.  3  is  in  the  Box.  Gazette  36:161-186,  280-302,  349-367,  4*9-455.  1903. 

81 


I 


BOTANICAL  OAZHrTE 


(ai'uuit 


An  li»  prcviiius  litcraturt-  of  thi(»  partiiular  bubjtit,  thcrt'  i.s 
none.  In  1886  Dr.  (J.  U.  Hay  made  a  colliiiion  i>f  Miscou  plants 
for  the  Ucologiial  Survey  of  Canada,  but  no  aicounl  of  them 
was  ever  published,  and  no  other  liotanisl  has  heretofore  been  tin 
the  island.  In  many  resfx  its,  however,  as  the  reader  will  observe, 
the  vegetation  of  this  bea.  h  plain  resembles  closely  the  vegetation 


no.  I.— Outline  map  of  part  of  the  Gulf  of  St.  Lawrence,  to  show  the  geographical 
position  of  Miscou  Island. 

of  the  sand  beaches  and  dunes  of  Lake  Michigan  as  described  in 
CowLKs's  well-known  memoir,'  and  many  of  the  conclusion^  of 
that  work  are  also  applicable  here. 

Grande  Plaine  extends  along  the  v.est  side  of  Miscou.    Begin- 
ning on  the  south  at  Eel  Brook  (see  the  accompanying  map,  fig.  2), 

>B0T.  GAMTTE  37:95-117,  167-JOJ,  281-308,  361-391.    1899. 


looft)  C.tNOXa-XASrF.XT  FOKEST  OF  UlSCOf  BF.ACll 


H 


where  it  is  but  a  f-w  yards  wide,   it   rapidly  >  r..adtns  northward 

until    it    reaches    Mune    half    a    mile    ;i  ross,    and    ihen    narr  -wi» 

URain  tuward-H  its  mtrthern 

end,    which    is    also    the  Btty  CAnitur 

northernmost  p.»int  ol  ihi 

island.     Though    nearly 

level  as  a   whole,   .!  is  by 

no  means  t^at,  for  it  is  com. 

IK)8cd  of  a  series  of   ap- 
proximately concentric  dune 

beaches,  which,  two  or  three 

in  numlK-rat  Kel  BnK)l(,  in 

crease  to  over  forty  opposite 

Lac  Krye.     In  height  these 

dune  be.rhcs   vary  from 

tv;o  to  five  or  six  or  even 

seven  feet  (o.6-a"')  and   in 

breadth  from  eight  or  ten 

up  to  forty  or  fifty  paces. 

At   iis   widest  part,  wnich 

'•omprises   s»)me    thirty  or 

more  of  the  beaches,  new 
ones  are  plainly  being 
ra-^idly  added,  while  at  its 
nortnern  end  the  entire 
plain  is  being  washed  away 
by  the  sea,  which  is  cutting 
sha.7>ly  across  the  ends  of 
the  old  Ijeachc  Aiiout 
two -fifths  of  the  plain,  in- 
cluding the  older  parts  n;xt 
the  upland,  are  lorested; 
about  two-fifths,  incluriinjr, 
all  the  outer  and  mv.er 
parts,  are  open,  cloi..ed  i  nly 

by  the  waving  beach  grass; 

the  intermediate  zone,  a  sr  .11  one-fifth  of  the  area  of  the  plain,  is  a 


»4 


BOTASICAL  liAZETTK 


(AI'OVIT 


trutiMlion  iimc  whtn-  the-  f-.rusl  is  pushinj?  iu  advamc  into  the  upcn 
Kruiiml.     Thf  mntU  of  f..rn»ali..n  of  this  nmarkahk'  |(lain,  invulv 
itif,'  ihf  an.nnaly  nf   fXliriMvc   land  huildinu  <,n   a  -inkinn  ^'mM,   I 
ha\i    ili's(  riU'd  w.miwhat   fully  tlMwhiri'. '      Uriilly,  ihf  fa(W  arr 
ihtM'.     As    tin-    risull    ot    pcculiariliijt    c-f    iho   ti.i)<.Kra()hy,  wiml, 
anil    liiUs   .if   this   rif^iim,    ihcro   i-.   f..rmcil  .m  iho  shallow  north- 
wi-sti'rn  Hiilf  ..f  Mihc.ii  a  kimi  of  (jnat  .cMy  iti  whirh  all  mo\at)lf 
nialtrials,  sand  ami  uravil  from  thf  wtiir  of  the  rapitlly  crumhIinK 
adjaifnt    coasts,    as    will    as    driftwood,    watirwcrds,    and    othtr 
tlolsam,  ofini  from  a  ^riat  distami-,  find  to  (olkii,  and  ihincf 
an-  driven  ii>horf  by  the  prevailing  wisltrly  wiruU.     I'ornurly  the 
island  ixlinded  fartlur  north  than  n-.w,  tarrying  wilh  it  Ix.th  i-ddy 
and  jilain;  but   ihf  >r<"<ral  subsidi.i.f  .k lively  in  progress  in  this 
region  has  tarried  ii>  low  northerly  i  nd  beneath  the  sea,  thus  fori  inj,' 
the  eddy  and  the  adompanyinjj  plain  buildin)?  >,'radually  southward. 
The  northern  end  of  (Irande  I'laine  tixlay  is  Ininf?  rapidly  washed 
away  (com|)nre  map),  to  be  redeiK)sited  farther  south,  and  the  plain 
as  a  whole  is  thus  rolling  by  its  outer  margin  southwaril  along  the 
eoasl.     The   subsidence   of   the   lanil  has  pr.dured   another  effeet 
U|)(.n  the  plain,  and  one  of  lonsiderable  eonse<|uen(e  to  its  ve^e 
tat  ion,  namely,  its  inner  and  oliler  |)art  averages  somewhat  lower, 
that  is,  less  above  sea  level,  thiin  the  outer  and  newer  part,  thus  lead- 
ing to  a  settling  of  water  towards  the  older  inner  parts,  and  a  rela- 
tively higher  water  table  in  them.     That  we  have  here  a  Ix-aeh  plain, 
instead  of  a  series  )f  lofty  sand  dunes,  is  the  result  of  l\  •  fact,  appar- 
ently, that  the  dry  sand  of  the  beach  is  blown  rshore    o  faster  than 
the  Inach  grass  can  fix  it.     .At  l>oth  the  northern  and  s(,uthcrn  enis 
of   the   plain,  however,  there    is   some   ajjproach  to  a  building  of 
true,  though  low,  dunes. 

My  brief  study  of  the  vegetat-on  of  Orande  Plaine  was  entirely 
observational,  not  at  all  instrumental,  nor  do  any  meteorological 
or  other  exact  physical  data  for  this  region  exist.  Orande  Plaine 
lies  at  sea-level  in  latitude  48°,  Ix-side  a  shallow  sea,  warm  in  summer 
but  frozen  over  in  winter.  The  summer  climate  is  remarkably 
equable,  of  a  temperature  most  comfortable  for  man,  with  no  fogs 
and  but  little  iloudy  weather.     The  rainfall  must  lie  not  far  from 

tBull.  Nat.  Hist.  S<h-.  N.  It.  No.  24:455,    11,06. 


I.K)'ll 


(;,i.V(>.v<;-,v.i.vt>:.vy  tnnKsr  of-  mi. 


Mxn   HI  t<  // 


45  iiKlus.     Hiiivy  wind;,  irinn  ilu'  wi>i  prtsiiil 
JMiil  is  of  imn-  or  ;rt/,  nanil  tk-riwil  in.m  tb 


in  siininu  r. 


Ilu 


ifcroii!*  Mimlstnms  <>f  ih 


»•  W(iii  II  ihf  ){ray  larU. 


I    ii'Kion,  this  naml  havi 


nx.  '  f 


iht 


usual  rdalion.-.  Id  waur  Mipph,  minira)  miinVnN,  .1..  N,,  .ili.r 
»|>i-iial  faiti.rs  with  .1  UaririK  u|n  n  \V  v.xiiaiioii  ,,,,|,t.ar  (..  Ik' 
promint-nt. 

Wc  iiirii  now  to  ton-.i(liT  the  Mnitalion.  AlihoiiKh  it  |.ri>tniH 
rwry  Krudatjon  fr.  m  Inin.l.lf  lurh-  .  |  ilu'  (.|Hn  Ua.  h  i<.  ilu'  (knsfM 
W(hkI,s,  nivirlhcless  ihi-  lyi-  Ih,  (  ,  a.tiihl.nuil  to  riT.Kni/i.  an<l 
Ihi-  siRTih  to  rk'>ignali',  (.rtaiii  .Klmilr  M'^ilational  rr>;i.ins.  'I'ln'M' 
ri-priHnt  the  mn.li..,  „r  .limaxis,  as  it  w.ri',  in  thi>  MKitati..p  ,  urM' 
-Ihi'  parts  whiih  ixhibit  a  iliMin.  live  ('lara.tiT  in  the  ,,hvsio«- 
nomy  of  thi-  whole  Ihiy  a  v  the  iVllosinj;;  (i)  ihr  new  Ina.h, 
(2)  ihi'  ^-rass  plain,  (.;)  the  swaks,  (4)  ilu-  samlv  w,m«I>,  (,)  ihv 
cU>8f<l  woodx. 

THK    NKW    BK.ACir. 

The  iharaitfri.stic  opm,  or  mw,  Ixiuh  of  Crande  I'laim.  the 
kinil  whkh  Ik-sI  illustratts  the  m.  dc    .f  ^rr„wth  .f  the  plain,  is  to 
Ix-   found  op|,<.sitf  its  mid.i:..  and        .adist    part;  for  towards  the 
northern  and  southern  en.ls  its  sin     >.re  is  niorliiied  in-  Imal  c  ndi 
lions  of  erosion  and  <lune  building.     Outsi.lc  .  f  all  is  a  hr.ad  sloping 
inter-lidal  beach  of  pure  sand  with(  ut  vegetation  itiii.  3).     Alx.ve 
it  is  the  ..arrow  band  Ixtween  onlinary  and  exirer      high  tides, 
from  which  the  drying  .sand  i.s  being  driven  landward        ,he  winds- 
it  1.S  aI,so  vegetati(  nless,  ,  r  .vilh  but  stragglers  fn.m  the  ..,,|ar  U-ach.' 
Finally,  there  is  that   broad  shelf,  very  well  shown  in  the  atc(  m- 
panying  photograph  (fig. .,),  reached  only  by  the  very  highest  tides, 
comp<sed  of  fine  t.uartz  sand,   intermixed  with  some  gravel  and 
occasional  flat  cobble.;;   it  is  (overed  with  scattered  <iriftwood  among 
and  over  which  the  dry  .sand  i.s  l)eing  f„rever  driven,  shifted,  and 
piled.    Thus  the  new  beach  offers  a  barren  habitat  to  plants,  for 
it  has  a  mineral-poor  soil,  drenched  <  ften  by  salt,  fcrever  shifting, 
and  c.xi:M).sed  to  the  unbroken  f(,rce  of  fretjuent  heavy  winds,     'i-hc 
vegclalioii  is  plainly  resp<>nsive  to  these  conditions.     It  is  extremely 
scanty,    the    plants    growing    widely    isolated,  while   manv   square 
yards  do  not  show  any  vegetation  at  all.     Thus  c.  mpetiti.in  among 
the  plants  seems  not  to  exist,  and  the  struggle  is  wholly  with  the 


86 


BOTANICAL  CAZETTh 


[august 


physical   environment.    The   most   characteristic   plant    by   far   is 
the  small,  radiate-decumbent,  succulent,  annual  saltwort,   Salsota 


y  "/i\^^''  "nir  //||„,'' 
'///n"y//ni>"%//iiii>'';„„. 

'/llllV7/(lll"'"///lillW"''«|,>'' 

mm 


2 

o 

e  ^ 
.2  2 


a  I 

U.   it; 


Kali,  which  occurs  but  rarely  and  for  the  most  part  in  the  lee  of 


1906] 


OANONG-XASCJlXr  FOREST  OF  MISCOC  liE.lCll 


87 


some  shelter,  such  as  a  hollow  or  large  piece  cf  (Iriftwood.  Xcxt 
in  abundance,  though  but  scarce,  is  the  little  fleshy,  nsette-like, 
annual  sea  rocket,  Cakile  edentula  (C.  amcricana).  Third  in  abun- 
dance is  the  low  haio-n  sctte,  perennial  sea  lungwort,  Mtrlinshi 
marUima,  here  seemingly  growing  as  an  annual,  also  m(  sllv  in 
places  of  some  shelter.  Rarely,  and  then  only  in  a  sheltered  ],<  si'tion, 
occur  tiny  radiate-creeping  plants  of  the  beach  pea,  Luthyrm  marl 
timus,  growing  apparently  only  as  an  annual,  and  sometimes  show- 
mg  a   marked  difference   in   the  windward-creeping  and   leeward- 


FlG.  4.— lypical  open,  or  new,  beach,  l.wking  north;  among  the  drift«i»(l  occur 
scattered  tufts  of  saltwort  and  Ijeach  grass. 

creeping  shoots  on  the  same  plant,  the  former  being  much  shorter 
and  smaller-leaved.  Even  rarer  is  the  radiate-creeping,  small- 
Ifaved,  halo-scurfy  annual,  Atriplex  patiila  haslala.  Here  and 
there,  however,  especially  in  sheltered  places,  arise  the  tufted  culms 
of  the  beach  grass,  Ammophila  arenaria,  the  characteristic  sand- 
binding  perennial  of  the  dune  beaches  next  to  be  considered,  here 
seemingly  growing  from  seed.  I  was  able  to  find  no  other  plants 
on^  the  new  beach.  Thus  we  see  that  this  vegetation  is  distinctly 
adjusted  to  the  physical  conditions,  for  it  is  (,f  great  jjaucitv,  of  small 
and  slow  growth,  annually  renewed,  closely  ground-apprcssed,  and 
strongly  xerophytic. 


88 


BOTANICAL  GAZETTE 


[Arousi 


THE  GRASS  PLAIN. 

Of  atlV^'  ';"'  "!  "'^""  ^''^  '^^'■"^  ''^'^  ^'^"d  plain,  composed 
!l  fhT  "T  °^/PP'--™-'<^ly  parallel  dune  beach  s.  forming 
smoothly  swelhng  ridges  and  hollows  r.f  elevations  and  breadth! 
already  descnbed.  Every  dune  beach,  I  believe,  originates  wU 
a  core  of  dnftwood.    As  the  tidal  beach  is  built  outwards  byle 

par  until  rif'  '"''"''''  '^™'"""  '"  ^*«  ""  ''^  "PPe-oM 
part,  until  finally  some  unusual  combination  of  great  winds  with 

high  t.dcs  sweeps  it  up  beyond  reach  of  further  disturbance     Thn 

the  dnvmg  sand  from  the  beach  is  caught  among  it;    the  beach 

grass  gams  a  foothold  in  the  shehered  places,  spreads,  and  cheks 

he  further  movement  of  this  sand.    Then  more  sa;d  is  dr'en 

shoreward,  and  it  grows  into  a  low  dune  which  is  fixed  bv  the  beac^ 

ST-fT  ^'  r  \"""    ''''  ""'*  '^  ^^-'-d  -ly  -hen' a  new  1  ne 
of  dnftwood  has  been  formed  outside  and  begins  to  stop  the  sand 
or  its  own  growth.    The  resultant  dune  beach  offers  severe  cond' 
ttons  for  plant  he,  for  its  surface  is  swept,  especiallv  on  the  summ  t 
and  wmdward  slope,  by  heavy  winds;  it  is  heated 'intensely  bv  the 
sun;  >t  .s  readdy  movable;  and  it  forms  a  soil  extremely  p^r  in 
mmeral  nutnents.^    It  lacks  the  salt  of  the  newer  beaches.'ho:e  -er! 
or  this  ,s  soon  removed  by  the  rain;  and  it  possesses  an  ampl 
supply  of  moisture  a  foot  or  two  beneath  the  surface  for  the  sud^v 
brought  by  the  rain  drains  but  slowly  away,  owing  to  the  Lw  ^ad  em 
of  the  water-table.    These  conditions,  especially  at  their  extreme 
on  the  summits  and  windward  slopes  of  the  Lch  dunes.        enlrTd 
by  practically  but   a  single  plant-the  herbaceous-perennial    Tb 
terranean-creeping   xero-culmed,  deep-rooting  beach^rass.  iZo- 

fn  t""'  7'"^'  """""■   "''•^""^  '-y  --I-titor  ;hatever 
m  open  sea  te^d  tussocks,  only  partially  coverinrthe  ground    ^ 
wel  shown  in  fig.  5,  and  in  closer  view  in  figs.  6  and  7.    iZLZ 
that  this  grass  is  of  considerable  economic  value  to  the  neighC 
mg  farmers,  who  cut  it  and  haul  it  for  hay.  and  whose  catt^'grte 

oTthe  Jut^'  btaT""  "•'"'  ^''  ^^"^^^  ^"  '-^'-  -po- 
of the  outer  beaches,  permitting  them  to  be  irregularly  cut  by  the 

wind.    It  IS  for  this  reason,  I  have  no  doubt,  the  newer  outer  beaches 

^^^^^As  indicated   by   Ke.k..Vs  receni   studies:    Box.   Gazkxtk   37:4.^436. 


.906]  GAmxC-NASCnXT  FOREST  OF  MISCOU  BEACH  gg 

are  SO  much  more  irregular  in  their  various  character,  than  the 
older  inner  beaches,  which  antedate  the  advent  of  man 

esped  li?£hfnd'^"f  '"?  '"  ""  ^""P^"'"^'  '^  ^'^-^•^  ^  shelter. 
oH ^e    nit         T-  uT"""'  P''""'"'"^  ''''  «^"^-"^  "^  -  ""-"be 
1  InHr^        '  '  ^''''''''"'  ^"'■'"  ''"'  ^"  '-nsignificant  part  of 

Inothe  m"':'''""'  ''"'  "''^'^  ^"  -^^•»>'  -P-ated  from  o  e 
Catr'smT  rr""'  "'  ''"^'  P'^^^P^'  '-^  '"^^  beach  sedge 
aZ  t  t  \  ^"""^^  '"  '^^"^^^'^  '"««"^ks  here  and  there 
among  the  beach  grass,  and  it  is  indeed  the  only  other  plant  Jhth 


theblrchS'''"'  "'^  """■  """"'^  "°"'^   P"*^"^""-^  "°  P"^"'  -""e  except 

seems  at  home  in  this  situation.  The  remainder  of  the  plants.  aU 
sparsely  or  rarely  represented,  are  obviously  stragglers  from  the 
most  diverse  habitats,  many  of  them  quite^unex^ct  d  tTdem 
Lh  anVt'eT"H  ''^"^' ^^"^^  ^'^-^  stra^  in  from  the 
on  i  t  L  u  IT  ''  ^'"^  '°'"^^'''^'  """"^  flourishing  than 
on  the  open  beach.  Then  there  are  greatly  dwarfed  individuals 
of  cjrtam  ubrquatous  forms  able  to  endure  a  wide  range  of  phyid 

hist    S,-r     "■"         :  °"'"  ""^"  ''  '""^  P'^'"'  ''^^  Canada 
thistle,    Cmcus   arvens^s;  the    sow    thistle.    Sonchus   arvensis;  the 


90 


BOTAXICAI.  GAZETTE 


[august 


field  sorrel,  Rumex  Acetosella,  in  dwarfcd-rosctte,  very  red  forms; 
the  evening  primrose,  Oenothera  biennis;  and  the  moonwort,  Botry- 
chium  ternatum  intermedium.    There  are  also  some  forms  usually 
characteristic  of  rather  a  dry  habitat,  such  as  the  pearly  everlasting. 
Anaphalis  margaritacea,  and  a  western  yarrow,  Achillea  lanulosa. 
In  addition  there  are  others,  generally  in  more  sheltered  spots  and 
also  greatly  dwarfed,  which  usually  prefer  a  moister  habitat,  such 
as  the  two  westc  n  roses,  Rosa  acicularis  (R.  Sayi,  R.  Engelmanni) 
and  Rosa  lucida;  a  western  violet,  Viola  adunca;  a  stitchwort,  Stel- 
laria  long! pes   laeta;   a  silver  weed,   Potentilla  Anserina  concolor; 
the  stellate   false  Solomon's    seal,  Smilacina   stellata;  and   one  of 
the  vetches,   Vicia  Cracca.'    The  great  diversity  of  natural  habit 
()f  these  plants,   their  scanty  and  irregular  occurrence,  and  their 
dwarfed  size  and  rosette-forming  tendency  all  unite  to  show  that 
none  of  them  are  here  at  home.    Obviously  they  are  the  ones  which, 
of  all  the  many  kinds  which  must  be  brought  to  this  plain  by  natural 
modes  of  dissemination,  are  sufficiently  tolerant  physiologically  to 
be  able  to  germinate  „..der,  and  then  to  withstand,  these  extreme 
physical  conditions,  eking  out  here  a  starved  and  precarious  exist- 
ence.   The  conditions  for  germination   upon   the  sand   must   be 
extremely  severe,  and  it  is  very  likely  that  other  kinds  of  plants 
could  exist  here  as  adults,  could  their  seeds  develop;  and  further 
It  IS  probable  that  the  individuals  which  do  exist  on  the  plain  are 
those  whose  seeds   happened   to  fall  in  especially  favorable  spots, 
or  became  properly  buried  by  the  moving  sand.     Else  why  are 
they  so  few?    The  universal  dwarfing  is  due  in  all  likelihood  not 
to  the  heat  and  dryness  of  the  surface,  nor  to  any  salt  content 
in  the  soil,  and  certainly  not  to  a  scarcity  cf  soil  water,  but  to 
the  paucity  of  mineral  nutrients  in  the  sand.    This  is  in  harmony 
with   another   feature    they   mostly   show   in   common-very   deep 
and,   I   think,   much-branching  roots.    The  fact  that   they  come 

5  The  following  Grande  Plainc  plants  appear  to  he  new  to  the  flora  of  New  Bruns- 
wick: .!./:,/,«  lanulosa.  Viola  adunca,  Rosa  acicularis,  Stdlaria  longipts  laeta,  and 
PolenUlla  Ansenna  concolor.  Certain  others  are  new  in  name,  the  species  having 
been  recently  more  exactly  defined  and  segregated:  .ilnus  mMs,  Myrica  carolin- 
rns,s,Vaccnuum  VUis-Idaea  minor.  Others  are  new  in  name  txrause  made  to  con- 
form to  the  riiles  of  the  Vienna  Congress,  but  in  these  cases  the  names  of  Gray's  Man- 
ual,  6th  edition,  have  Ixen  given  in  brackets. 


«9o6]         CANONG-NASCRNT  FOREST  OF  UtSCOV  BEACH  91 

from  such  a  diversity  of  natural  habitats,  and  yet  hve  in  this  peculiar 
situation  upon  an  equal  f(K)tinp,  shows  how  far  we  are  from  under- 
standing the  real  bases  of  physiological  adaptation,  and  further 
shows  that  in  the  study  of  the  physiological  life-histories  of  plants 
lies  the  most  important  and  attractive  field  for  the  ecologist  of  the 
near  future. 

So  much  for  the  exposed  parts  of  the  dune  beaches.  But  in 
addition  they  offer,  upon  their  inner  or  leeward  slopes  and  in  the 
hollows,  situations  more  sheltererl,  not  so  much  from  the  sun,  since 


Fic.  6.— Typical  hollow  between  outer  dune  beaches;    the  tall  grass  is  all  beach 
grass,  but  the  small  plants  among  it  are  the  common  strawberr)-. 

their  average  course  is  nearly  north  and  south,  but  from  the  westerly 
winds.  The  older  inner  dune  beaches  also  are  protected  to  some 
extent  by  the  newer  outer  ones,  as  well  as  by  their  slightly  lower 
average  level.  The  diiference  between  leeward  slope  and  hollow  is  not 
simply  one  of  degree  of  shelter,  however,  but  also  of  physical  condi- 
tions, for  the  hollow  is  much  nearer  the  source  of  water  supply, 
the  free  table  of  which  is  not  over  a  foot  or  two  beneath  the  surface! 
In  consequence  of  these  differences  we  can  recognize  three  distinct 
phases  of  vegetation:  prst,  a  larger  development  on   the  leeward 


9» 


BOTANICAL  GAZETTE 


[aitoust 


li- 


slopcs  of  plants  which  are  small  and  rare  on  the  outer  slopes;  second, 
a  distinctive  vegetation  of  the  hollows;  and,  third,  a  distinctive 
vegetation  of  the  inner  slo})es. 

As  to  the  first  phase,  it  is  enough  to  note  that  several  plants, 
in  particular  the  pearly  everlasting  and  the  wild  roses,  small,  scarce, 
and  scattered  on  the  outer  slopes,  become  larger,  frequent,  and 
even  patch-forming  on  the  inner;  and  this  is  true  alsD  in  less  degree 
of  other  species.    The  beach  grass  persists  in  all  situations. 

The  second  phase  is  the  vegetation  of  the  hollows.  The  very 
first  plant  to  appear  in  this  situation,  and  that  too  near  the  outer 
beaches,  is  always,  strangely  enougii,  the  common  wild  strawberry, 
Fragaria  virginiana,  apparently  of  normal  size  and  form,  seem- 
ingly quite  at  home,  and  spreading  abundantly  by  runners,  so  that 
it  forms  considerable  patches.  The  appearance  of  the  nascent 
patches  is  wel'  shown  in  fig.  6.  As  soon  as  the  patches  reach  an 
appreciable  density,  such  that  they  afford  a  cover  to  the  ground,  then 
turf-forminf?  grasses,  of  which  the  first  is  the  red  fescue,  Festuca 
rubra,  appear  and  initiate  the  turf-formation  which  is  so  important 
a  feature  of  the  swales  to  be  described  below.  The  strawberry, 
of  course,  is  one  of  the  most  tolerant,  and  hence  ubiquitous,  herbs 
of  our  flora,  and  its  situation  here  is  partially  explained  by  the  near- 
ness of  the  abundant  water  supply.  Yet  it  is  surprising  to  find 
it  taking  so  important  a  part  in  a  vegetation  in  so  peculiar  a  position. 

The  third  phase  of  this  vegetation  is  that  characteristic  of  the 
sheltered  slopes.  First  of  such  plants  to  appear,  and  the  most 
common  and  characteristic,  is  the  dwarf  creeping  juniper,  Junip- 
erus  Sabina  procumbens,  of  which  single  plants  originate  just  below 
the  beach  dune  crests,  and  creep  radiating,  more  to  leeward  than 
to  windward,  in  a  close  dense  mat  covering  many  square  feet.  A 
young  plant  is  show"  in  fig.  y,  in  characteristic  form  and  position. 
On  the  inner  beaches  ihese  plants  occur  upon  the  oaier  as  well 
as  the  inner  slopes,  and  the  shelter  of  the  mats  thus  formed  affords 
in  reality  the  principal  starting-point  for  the  development  of  other 
plants  which  lead  gradually  to  the  development  of  the  forest,  as 
will  be  noted  under  the  transition  vegetation.  In  a  similar  situation, 
but  independently,  arise  patches  of  two  other  characteristic 
plants,  a  bright-green,  leathcry-lea\ed,  tufted  shrub,  the  wax  berry. 


iQOft]         CA!VO\'a—NASCEXT  FOREST  OF  MISCOU  BEACH 


93 


Myrica  carolitunsis,  which  comes  to  form  dense  discoid  (sometimes 
almost  fairy-ring  like)  masses  (.n  the  crests  and  inner  slopes;  and 
the  less  frequent,  low,  dense-tufted,  white  hairy  shrub,  Hudsonia 
tomenlosa,  in  irregular  close  patches.  All  (.f  these  plants  are  pro- 
nounced xerophytes,  which  amply  ex;>lains  their  ability  to  live  in 
this  situation,  and  even  their  preference  for  the  somewhat  drier 
upper  slopes  of  the  dune  beaches.  Their  xen.philism,  in  common 
with  that  of  many  other  evergreen  sand  plants,  is,  as  I  guess  it,  an 


8W!i 


''^N  't^^;^**^ 


t  ,i«(»*r'ir*.. 


■^^^ 


Fig.  7. — Typical  upper  inner  slope  of  a  dune  beach;  the  grass  is  beach  grass,  but 
in  the  center  is  a  typical  plant  of  dwarf  creeping  juniper. 

adaptation  to  the  physiological  dryness  which  results  when,  as 
must  often  be  the  case  in  spring  and  fall,  the  ground  water  is  of 
low  temperature  and  hence  slowly  absorbed,  while  the  leaves  are 
exposed  to  high  tianspiration  conditicns  from  the  bright  sun,  heat 
reflected  from  the  .sand,  and  high  winds.*  The  juniper,  while 
perfectly  at  home  here,  apparently  is  so  cnly  thrcugh  coincidence, 
for  its  original  habitat  is  seemingly  dry  reeky  hills.    But  the  other 

*  This  principle,  which  from  its  discoverers  we  may  call  the  KtHLUAN-GoEBEL 
principle,  seems  to  me  deserving  of  much  more  recognition  than  our  students  are 
inclined  to  give  it.     At  least  it  calls  for  careful  experimental  investigation. 


94 


BOTANICAL  GAZETTE 


[auovst 


i 


i|' 


two,  the  waxberry  and  the  Huclsonia,  are  characteristic  of  just  this 
situation,  in  and  to  vhich  they  have  apparently  been  adaplively 
developed.  Towards  the  inner  dune  Ix'aches  another  low  shrub 
comes  in  on  the  slopes,  though  dwarfed  and  not  abundant,  the 
common  blueberry,  ,ccinium  pennsylvanicum;  it  is  evidently 
not  here  at  home,  bu  Us  somewhat  xerophytic  habit  permits  it  to 
exist.  As  these  various  plants  grow  older  and  extend  their  patches, 
they  run  together  more  or  less,  sometimes  two,  sometimes  three, 
and  even  all  four.  Later  others  are  addi'  to  them,  initiating  the 
juniper  mats  and  the  wcwds  carpet,  later  to  be  considered. 

The  contrast  lx;tween  the  vegetation  of  the  outer  and  the  shel- 
tered slopes  of  the  dune  beaches  comes  out  with  striking  clearness 
a  few  hundred  yards  north  of  Eel  BrcHik,  where  it  happens  the  entire 
plaii.  is  very  narrow,  and  sIcjIjcs  in  both  directio.  s  from  a  central 
higher  crest.  Outside  of  this  can  be  seen  only  the  l)cach  grass  ami 
its  accompanying  forms  as  listed  above,  while  inside  the  various 
xerophytic  shrubs  show  to  great  perfection. 

THE  SWALES. 

Between  the  open  grass  plain  and  the  woods  occurs  a  transition 
zone  marked  n')t  only  by  an  intermediate  vegciation  but  also  by 
distinctive  physical  features  as  well.  First  of  all  it  is  characterized 
by  the  presence  of  several  great  lurf-carpeted  and  tree-bordered 
swales,  morphologically  hollows  l  'ween  the  dune  beaches  which 
here  spread  much  farther  apart  than  usual.  They  are  well  shown 
in  figs.  8,  10,  II.  They  are  best  developed  in  the  widest  part  of  the 
plain,  hardly  cccurring  towards  its  southern  or  northern  ends,  and 
outside  of  them  runs  a  line  of  higher  dune  beaches,  which  indeed 
can  be  traced  through  most  or  all  the  length  of  the  plain  (fig.  z). 
The  swales  are  narrow  southward,  but  broaden  northward,  deepening 
as  they  go,  until  in  some  cases  they  dip  beneath  the  water-table 
(thus  exhibiting  pools),  after  which  they  rapidly  narrow  and  rise 
to  disappear  northward.  Again,  the  trees  of  this  zone,  occurring 
always  along  the  slopes  of  the  dune  beaches,  do  not  exhibit  a  transi- 
tion of  size  and  age  to  those  of  the  sandy  woods,  but  are  always 
so  much  smaller  and  younger  as  to  be  sharply  marked  oflf  from  them, 
the  case  shown  in  fig.  lo  being  very  exceptional,  and  that  of  fig.  8 


«9o6)  GAXOXa-ffASCEXr  FOKEsr  OF  MlSCOt'  BF.iCIl  95 

more  typical.  Again,  the  transition  fn.rn  the  bnmd  swales  t..  the 
beaches  of  the  sandy  vmkIs  is  m,.st  abrupt,  f„r  the  latter  are  regular 
narrow  close  together  with  scarcely  any  hollows  between,  and 
also  exhibit  a  curious  barre-ness  on  their  summits  in  marked  en- 
tra.st  to  the  better-dothed  summits  farther  out  (cmpare  /igs.  H 
and  13)  Unfortunately  the  full  importance  of  these  features  did 
no  stnkc  me  m  time  for  a  study  ,  f  ,hcm  on  the  ground,  but  ,uch 
data  as  I  possess  in  n<.tcs  and  maps  lead  mc  to  believe  that  the  swales 
arc  much  newer  in  origin  than  the  beaches  imme.liately  inside  them', 
ard  that  they  mark  the  transition  from  an  ,  Idcr  series  <,f  Jx-aches 
which  formed  part  of  the  original  Grande  Plaine  extending  far  to 


withS'Ll^mT^'''")'/™"''"""  """'■  '""'''"«  """''•  "•""^■'"8  <»  »*»'<'  °n  the  right, 
with  Its  sharp  Unc  of  tmns.Uon  to  the  woods;    the  trees  are  all  white  spruce. 

the'northof  the  present  i.land,  and  a  newer  series  formed  by  the 
rolhng  of  the  plain  down  the  coast,  as  described  earlier  in  this  paper. 
Al  the  facts  I  possess  both  as  to  geography  and  vegetation  are  con- 
sistent with  this  view.' 

Aside  from  the  question  of  age,  the  swr'e  zone  differs  physically 
from  the  grass  plain  by  its  greater  shelter  from  the  west  winds,  its 
lower  level  and  greater  nearness  to  the  water-table,  a  probable 
increase  of  mineral  nutrients  derived  from  decaying  driftwood  and 
diffusion  from  the  upland,  and  some  slight  accumulation  of  humus. 

T  And  it  is  sustained  by  the  tradition  of  the  R-sidents  who  say  that  the  plain  has 

X  thi:  Tu^d^r  T"  """"^  """°^'  *''•""  "'^  memory'^f  ^^su" 
24  4?6.   ,906  '^  ^'■"  '""'^  '""^'  '"  ''""•  ^'''-  "'^'-  S"^-  N.  B.  No. 


BOTANICAL  GAZETTE 


[^vavn 


becoming  the  bn,ad  expanse  of  mea.low  turf  of  the  .wales    an.l 

the  jumper  mats  extemling  anM\y  with  the  a.Miti,  „    f 

white  «nni.-.. .      c     !•    •    '"'^     ""'"'•"'''"'""  <'f  many  young 

2    ;-7       r         Tr*  '"""  ""^'  """"'^•^'  "^"'  'here  re.ult  gla  les 

"u  inX  t  '^      ' """  '■'"""■"«  "'^"^•'''  "-^  ^"-■"  -p^^i""y 

..  this  plant,  or  else  variously  unite.!  an.l  cmbined  wi-h  patches 
..f  waxberp:.  Hu.Isonia,  an.l  bluefxTry,  exten.l  greatly  in  .n'm  t  " 
a.verjng  the  crests  as  well  as  the  slo,x.s  of  the  .lunc  Ua  .7  i 
thus  form  a  w,KKly  net  in  the  shelter  of  which  several  otheLms 

especially  the  beach  ^., .ss,   ,H-arly  everlasting,   and  yart.w     The 
new  forms  which  a,,.ar  .re,  first  of  all.  the  common  cn^wl,  rry  E. 
M-  «.^r«.  and  the  rock  cranberry,  Vacciniun,  Vi,i.I,aJ.^ 

aH  of  them  plants  characteristic  ..f  dry  upland  rocky  situations 
Less  frequent  are  the  little  gentian.  Genliana  Amarel/a  acutJZ 

t^^ corn  f:  •'%"'"-^ "^ ^^p^^'^"^ -" ^-^•^^•'^ 

Palustre  (so  much  more  highly  developed  in  the  woods)  another 
curious  mixture  of  xerophytic  and   hyZphytic  f.,n^rVe  ha" 

lro7f.r  d  "  T^'T'  '  '■"''  '^'^^-geneous  assem! 
Wage  of  forms  drawn  from  diverse  natural  habitats  all  the  wav 
from  r.,cky  hills  to  bogs.     They  do  not  exist   here    th  rctre  tn 

forms  of    he  flora  .>f  this  region  whose  adaptations  happen  to  fit 
he^  conditions,  or  wh.>se  range  of  physiological  ...leratioThappen 
o  be  great  enough  to  permit  endurance  of  the  conditions  here     Of 

helps  to  emphasize  h..w  large  an  element  of  accident  or  incident 


•906] 


ovi.vo.V(;-.v.i.vt/.;.vr  tonicsT  oh  .viscoi  bi:.u„ 


97 


then,   i,  in  „,,ap,a,i..„,  „„.,  ,,.,,  y^^^^.^^.   .,   .^  „,^^,  .^^^ 

u  .  matdy  pn.vc  ...  U-  a  ma..cr  ..f  ,hc  1.h..sc.  an.l  lar^c  rather  ,h«n 
ol  thf  ix ut  ami  mmutc. 

Finally    it   is  in  this  same  situation.  u|.,n   the  u,.,Kr  s1,.,h^  „f 

P  r<-a  ./«.    develop.     Stan.linK  in   ..,,en    f..rmatio, .   ,hey  d ,  no 
."terferc  wah  one  am.th.r's  «n.w,h,  and  in  o.n...,u.n,;  In-eome 


nunllr    f~'^yy''"'l  '"8^  J""'P«^  ■""'  ""  a  ^l"pe  an.l  crest  ot  ,  dune  Ix^a.h    with  a 
number  of  assocmted  planU  n..|e,l  in  the  text;   k  >king  s.,uth. 

™!/"'Tr'"'  ^•^^^•'\^>™'"^''"-'  '"  """ine  and  clothed  to  the 

Zrl'  I  7  'T^-  "'  '''"""■""  ""  ''""''^'  ^"^  ^'^'^  '^'^^  reason 
hat  the  shrubs  do  as  a  compromise  Ix-tween  the  greater  wetness 
...  the  hollows  and  the  greater  dryness  of  the  beach  summits.  This 
habtt  of  growmg  thus  upm  the  sloj^^s.  and  not  on  summits  .,r  h.,1- 
lows  has  a  most  important  effect  uiK.n  the  physiognomv  of  the 
veg^ta  ton  m  t  ,s  zone;  for  to  it  is  due  the  ...nness  of  the  swales, 
^th  thetr  regular  borders  of  trees,  and  as  well  the  openness  of  the 
beach  summits  m  the  sandy  woods  later  to  Ik.  noticed.  Toward 
the  sea  the  spruces  are  small  and  dense,  and  often  show,  as  in  fig  n 


BDT.i.MCAL  li.iZt.rTH 


(4U0IWT 


pronounud  wind  cffwls.  In  phucs  many  stwIlinK  'rif*  m»y  In- 
foumi,  thouKli  ihf  «liMri»)uti<)n  of  ihisc  U  curiously  irrcf{ular.  In 
one  plait-  only  did  I  find  any  .  thir  Irtr,  und  tha».  was  a  (tingle  .s|K.ti. 
mi-n  (.f  the  prinu's  pine,  Pinm  Punksiima. 

If  It  be  axkctl  why  the  white  spruce  is  the  first  tree  to  develop 
on  these  plains  instead  of  some  other  of  those  growing  on  the  upland 
near  by,  I  can  only  .>uy  that  an  answer  must  wail  until  we  know 
something  alxiut  the  physiology  of  the  wh"  iinue  and  of  other 
trees  of  the  viiinity. 

We  turn  next  to  the  swales,  those  long  ojKn  hollows  car|K-ted 
by  tt  close  turf,  ami  l)ordere<l  by  spruces.     The  general  apjK'arance 


Kio,  ;o.— Highly  developed  swalr,  looking  south,  on  the  left  U  the  edge  or  the 
Mil.!  woods  with  old  trtis,  and  on  the  right  a  lint  of  niu<h  younger  trees,  here  much 
largi  r  ihan  usual. 

of  the  turf  is  well  shown  -n  the  right  in  fig.  8,  and  extremely  well 
in  fig.  10,  which  shows  pe.  laps  the  f)est-devel()ped  of  all  the  swales. 
The  urf  is  a  gcKKl  deal  modified  in  vegetation  y  the  grazing  of 
cattle  and  horses,  yet  its  general  characters  sho.  plainly  enough. 
Originating  in  the  (,uter  hollows  with  the  strawberry,  as  already 
noted,  he  real  turf  Ix-gins  wit'i  the  red  fescue  grass,  Fesluca  rubra 
(F. ovina  rubra),  which  s(H)n  drives  (ut  the  strawljerry.  To  this,  as  it 
becomes  compact  in  the  inner  h')llows,  other  grasses  are  rapidly 
added,  especially  the  June  grass,  Poa  pratensis,  and  th?n  the  brown 
top,  Afiroslis  alba.  After  these  comes  a  rush  Junrus  Vaseyi,  and 
the  little  sedge,  Care.v  Oahri.     \-.ry  likely,  also,  there  are  other 


i.KJftj         ii.tSO.\a  -SASCKSr  tOKtCST  Of  MISCOf  HHM  n  i)Q 

Kra«M:»i  which,  owin^  to  my  !m{H  rlfvt  kni>wk-<|f(i-  of  thoM-  ki^>u|mi« 
I  ovi-rltMtkfil.  On  anil  iimnnn  thit»f  plants  iKtur  uthtrs,  anions 
whkh  I  have  colU'ilcd  the  followinn:  the  cytl»ri}(ht.  Euphrasia 
ameriiiiHti  lE.  i>t)'uinali\);  the  huKle  wird.  I.yiopus  uniflttrus  (L. 
I'irniniiinii  ;  a  liny  everlasting,  Anlinmirin  miHlioua;  a  }K'arIwi>rt, 
Sagina  profumhtm;  the  plantain,  Plunlano  mnjttr;  the  two  eonim4)n 
ein<|uefoils,  Polinlilla  mmrgka  and  Annrrina;  the  fall  dandelii  n, 
Ltontodon  aulumnair;  and  the  while  i lover,  Trijolium  rrhfns. 
These  forma,  in  tomnion  with  the  Kraskes,  are  all  greatly  d.       e<l 


Fir,.  It. — An  outer  swalr,  liviklng  north;  in  the  tenter  clum^  if  blue  flag;  on 
the  »l()|)e  on  the  left  white  spruce  und  ttaxbcrry;  on  th''  ripSt  u  a  low  depression  with 
a  tliiiket  of  (>>iplur  (the  white  spruce  among  it  Ix-ing  on  a  lixal  elevation). 

and  do!ived  from  diverse  habitats,  and  are  evidently  a  collection 
of  heterogeneous  stragglers  from  the  neigh lK)rhood,  held  together 
by  no  stronger  bond  than  ability  to  ckc  out  e.xistence  in  this  inhos- 
pitabl"  position.  The  majority  belong  to  somewhat  moist  places, 
and  they  find  an  ample  supply  of  water;  for  the  water-table  even 
in  the  driest  summer  is  within  a  f(x)t  of  the  surface,  and  of  the 
sweetest  water.  Evidently  it  is  not  dryness  which  stunts  the  forms, 
but  most  likely,  is  I  believe,  paucity  of  mineral  nutrients.  The 
tuif  represents  the  first  closed  formation  we  have  met  with,  and 


lOO 


BOTANICAL  GAZETTE 


[august 


competition  may  therefore  determine  some  of  its  minor  features, 
but  to  these  I  gave  no  attention. 

The  turf  reaches  its  climax  in  the  open  swales  like  those  shown 
by  fig.  10.  In  the  woods  it  disappears,  as  will  be  noted  under  the 
next  section;  but  towards  the  lower  levels,  especially  towards  the 
pools  of  standing  water,  it  gives  way  gradually,  by  definite  steps, 
to  an  assemblage  of  true  swamp  plants.  The  very  first  of  these 
to  appear  in  the  k  wcr  i)laces  in  the  swales  is  always  the  common 
blue  flag,  Iris  versicolor,  and  characteristic  scattered  clumps  of 
this  plant  may  be  seen  in  the  foreground  in  fig.  ii,  in  the  distance 


Fig,  I j.— Marshy  swale,  looking  south;  in  the  center  a  permanent  pool  with 
margin  irodden  by  catUe;  behind  it  are  cattails  and  rushes,  and  back  of  them  a  thicket 
of  poplar;  on  both  right  and  left  is  sweet  gale,  and  in  the  foreground  is  the  blue  flag. 

on  the  swale  in  fig.  8,  and  on  the  left  margin  of  the  swale  in  fig.  lo. 
Next  follows  always  the  sweet  gale,  Myrica  Gale,  and  after  that 
low  bushes  of  the  balsam  poplar,  Populus  balsamijera,  a  plant  which 
forms  very  dense  thickets  and  grows  larger  as  the  situation  is  more 
sheltered.  Finally  the  pools  of  standing  water  arc  reached,  and 
on  their  margin  occur  cat-tails,  rushes,  and  mare's  tail,  Hippuris 
vulgaris,  with  some  other  forms  which  I  have  not  attempted  espe- 
cially to  study.  The  plants  may  be  variously  combined  according 
to  local  circumstances,  but  a  \cr>-  typical  arrangement  is  shown  in 
fig.  12.    It  is  plain  that  we  are  dealing  here  simply  with  an  ordinary 


1906] 


GANONC— NASCENT  FOREST  OF  MISCOV  BEACH 


lOI 


swamp,  offenng  nothing  peculiar  unless  it  be  the  small  size  of 
some  of  the  plants,  notably  the  poplar.  But  these  places  develop 
yet  farther  in  time,  and  there  come  in  after  the  poplar  three  willows : 
ScUix  bdsamijera,  S.  lucida,  and  5.  Candida,  forming  very  dense 
thickets,  and  apparently  under  congenial  conditions.  Finally  comes 
in  the  alder,  which  appears  to  be  mostly  a  form  of  the  green  alder, 
Alnus  mollis,  giving  us  the  culmination  of  the  swale  thickets. 

THE  SANDY  WOODS. 
Inside  the  swale  zone,  through  almost  the  whole  length  of  the 
plain,  extends  a  narrow  zone,  only  some  four  or  five  dune  beaches 


Fig.  13. — Typical  sandy  woods,  just  inside  the  swales,  looking  north;  in  the  cen- 
ter a  dune  beach,  bearing  scanty  beach  grass  and  reindeer  lichen,  while  on  the  slopes 
are  small  juniper  mats  with  white  spruces. 

wide,  of  remarkable  sandy  woods,  whose  characters  are  well  shown 
'^y  1^&-  ^3-  Their  most  striking  feature  is  perhaps  the  relative  bare- 
ness of  the  tops  of  the  beaches,  which  remain  far  m(  re  clear  of  vege- 
tation than  do  most  of  the  beaches  outside  of  them ;  and  this  bare- 
ness, in  conjunction  with  the  presence  of  trees  on  the  slopes  and  in 
the  hollows,  gives  rise  to  curious  vistas  as  shown  by  the  photi  graph. 
The  bareness  must  have  some  physical  basis,  but  I  was  not  able 
to  discover  it.  These  dune  beaches,  further,  are  very  narrow,  low, 
and  regular,  with  hardly  any  true  hollows  between,  so  that  the  turf 


i02 


BOTANICAL  GAZETTE 


[aogvst 


from  the  swales  is  very  scanty,  almcst  wanting,  in  this  Z'  ne.  Further, 
the  trees,  all  of  them  white  spruces,  are  much  elder  than  those  of 
the  swale  zone,  the  transition  being  commonly  of  the  most  marked 
abruptness.  All  of  these  features  tend  to  emphasize  the  conclusion 
earlier  given,  that  there  is  an  abrupt  physical  difference  between 
the  beaches  of  these  woods  and  those  outside,  a  difference  which, 
I  feel  sure,  is  one  of  age.  The  position  of  the  zone  would  indicate 
that  it  possesses  more  favorable  physical  conditions  as  to  water, 
mineral  supply,  and  shelter  than  the  zone  outside,  with  which  the 
large  size  of  the  trees  is  in  agreement.  But  the  bigness  of  the  trees 
makes  the  barrenness  of  the  beaches  all  the  harder  to  explain.  In 
their  vegetation  the  sandy  woods  e.'hibit  three  divisions:  the  sparse, 
scattered  beach  grass  and  reindeer  lichen  (fig.  ij)  on  the  beach  crests 
already  mentioned;  a  few  and  sir  ill  patches  of  turf  which  can  hardly 
obtain  a  foothold  where  the  hollows  are  so  small;  and  the  juniper 
mats  in  the  slopes  and  hollows  with  their  well-grown  white  spruces. 
The  mats,  however,  are  no  longer  entirely  creeping,  for  the  junipers 
send  up  numerous  erect  shoots.  With  them  persist  several  of  the 
plants  from  the  transition  zone,  especially  the  rock  cranbeny,  the 
three-toothed  cinquefoil,  the  pearly  everlasting,  and  a  few  others. 
But  in  addition  new  forms  come  in,  especially  and  characteristically 
the  bearberry,  Arclostaphylos  Uva-ursi,  a  rocky-hill  plant,  here  creep- 
ing radiately  over  the  sand,  apparently  under  congenial  conditions. 
Beneath  the  shelter  of  the  trees  appear  some  plants  of  the  woods 
carpet  which  we  may  best  consider  under  the  next  section.  The  trees 
themselves  are  of  moderate  size,  rarely  if  ever  over  20  feet  in  height. 
A  fact  of  interest  about  the  juniper  mats,  applying  also  to  a  less 
degree  to  the  forest  mat  which  succeeds  it  in  the  closed  woods,  is 
its  very  slight  hold  upon  existence  on  the  sand,  for  where  teams 
cross  and  disturb  it,  the  entire  mat  dies  and  soon  disappears.  Such 
instability  shows  forcibly  how  hard  are  the  conditions  of  life  in  this 
situation,  and  how  narrow  the  margin  between  success  and  failure. 


THE    CLOSED    WOODS. 


The  climax  of  the  sand-plain  vegetation  is  reached  in  the  dense 
though  dwarfed  mixed  woods  extending  between  the  sandy  woods  and 
the  upland,    A  typical  view  of  the  closed  woods  is  shown  by  fig.  14. 


1906] 


GANOXG—XASCEXT  FOREST  OF  MtSCOU  BEACH 


103 


Physically  the  situation  is>  much  more  protected  than  the  zones  out- 
side of  it,  and,  lying  at  a  slill  lower  level,  it  has  a  moister  soil.  The 
soil,  however,  is  slill  of  sand,  though  it  contains  some  humus  from 
the  decaying  vegetation  and  must  derive  some  mineral  matter  by 
diffusion  and  drainage  from  the  upland.  Very  likely  also  the  sand 
is  shallower  here  than  farther  out  (fig.  2),  and  hence  some  influence 
of  the  minerals  of  the  underlying  s^il  may  be  felt,  while  in  places 
an  appreciable  enriching  of  the  soil  must  result  from  the  decay  of 


Fic.  14. — Typical  closed  woods,  chiefly  of  white  sprue  .  i  'it  with  some  deciduous 
trees  in  the  background;  the  closed  forest  carpet  shows  in  the  glade  of  the  left  fore- 
ground. 


the  bodies  of  the  walrus,  formerly  slain  here  in  great  numbers,  as 
manifest  by  their  semi-fossil  bones.**  These  additional  sources 
of  mineral  nutrients,  however,  by  no  means  furnish  a  supply  sufficient 
for  the  proper  growth  of  the  woods,  for  in  every  <'eature  they  exhibit 
marked  depauperation  as  compared  with  the  same  species  on  the 
neighboring  upland. 

In  relation  to  the  preceding  zone,  the  closed  woods  consist  essen- 
*  Described  more  fully  in  a  note  in  Bull.  Nat.  Hist.  Soc.  N.  B.  No.  24:463.  1906. 


104 


BOTANICAL  CAZETTU 


[kvovn 


tially  of  a  greater  development  of  the  juniper  mats,  which  unite 
to  form  a  complete  unbroken  carpet,  together  with  a  greater  devel- 
opment, both  in  number  and  size,  of  the  white  spruce  trees,  to  which 
are  added  some  deciduous  trees  and  shrubs.  And  where  the  hollows 
dip  lower  than  usual,  and  towards  the  upland  in  places,  this  forest 
merges  to  alder  and  cedar  swamp. 

We  consider  first  the  woods  carpet.  Morphologically  it  is  a 
direct  development  of  the  juniper  mats  of  the  outer  zones,  though 
but  little  juniper,  aside  from  occasional  erect  shoots,  is  left.  With 
it  persist  some  of  its  earlier  associates,  the  rock  cranberry,  three- 
leaved  cinquefoil,  some  grasses,  the  bearberry,  and  the  reindeer 
lichen,  varying  in  their  respective  development  according  to  situa- 
tion. To  these  are  now  added  dwarf  plants  of  the  bunchberry, 
Cornus  canadensis,  the  twin  flower,  IJnnaea  borealis  amer'.cana, 
Pyrola  chlorantha,  the  pipsissewa,  Chimaphila  umbellala,  and  an 
abundant  brown  moss,  which  has  been  identified  for  me  by  Mr. 
A.  J.  Grout  as  Aulacomnium  paluslre,  a  typical  swamp  moss. 
Upon  this  carpet  develop  a  few  larger  forms,  especially  the  abundant 
wild  sarsaparilla,  Aralia  nudicaulis,  the  gooseberry,  Ribes  oxy- 
acanihoides,  the  dwarf  raspberry,  Rubus  Iriflorus,  with  others  less 
conspicuous. 

We  consider  next  the  trees  of  these  woods.  First  in  importance 
and  size,  far  surpassing  all  others  in  both  respects,  is  the  white  spruce. 
It  attains  a  height  of  perhaps  7. 5*",  a  diameter  near  the  ground 
of  perhaps  45"",  and  it  exhibits  over  100  annual  rings,  though  per- 
haps some  may  be  much  older  than  those  I  counted,  which  were 
cut  by  the  residents  for  wood.  The  next  to  appear  is  the  balsam 
fir,  Abies  balsamea,  becoming  somewhat  abundant  and  character- 
ized by  a  spruce-like  arrangement  of  its  leaves  all  around  the  stems. 
Then  follow  the  red  maple,  Acer  rubrum,  the  aspen,  Populus  trem- 
ulaides,  the  paper  birch,  Belula  alba  papyrijera  (in  very  small  trees 
however),  and  the  mountain  ash,  Pyrus  americana;  while  the 
common  undershrubs  are  the  red  dogwood,  Cornus  slolonijera, 
and  the  black  alder.  Ilex  verticillata.  There  are  probably  some 
others,  but  these  I  believe  arc  all  that  are  notable. 

In  especially  low  places,  such  as  in  certain  hollows,  and  at  the 
contact  of  plain  and  upland,  the  conditions  verge  towards  those  of 


t{ 


igo6J 


CANONO-NASCENT  lOREST  OF  MISCOU  BEACH 


lOS 


a  swamp,  and  swamp  plants  apjK'ar — the  iris,  the  sweet  gale,  some 
mints,  species  of  Ga'.'um,  and  the  dewberry;  while  the  spruce  gives 
way  to  the  white  cedar.  Thuja  occidenlalis,  and  the  alder  beiomes 
abundant,  forming  a  dense  jungle.  But  this  is  of  less  interest  than 
the  vegetation  of  the  outer  zones,  and  hence  I  gave  it  little  study. 

Thus  it  appears  that  these  woods  present  no  features,  size  of 
the  plants  alone  excepted,  markedly  different  from  those  of  woods 
preponderatingly  coniferous  in  the  neighboring  upland,  and  they  arc 
evidently  tending  towards  the  typical  woods  of  this  region — the  mixed 
coniferous-deciduous  forest. 

We  have  thus  another  illustration  of  that  principle  so  important 
in  physiognomic  ecology,  that  vegetation,  no  matter  under  what 
immediate  physical  conditions  it  may  be,  is  always  tending  towards 
a  climax  type,  determined  primarily  by  climate. 


CONCLUSION. 

In  this  paper  I  have  tried  to  state  the  facts  about  the  vegetation 
of  a  somewhat  rcinarkable  place,  adding  thereto  some  ecological 
comment  whose  chief  value  is  to  illustrate  our  ignorance  of  that 
subject.  As  I  understand  it,  such  descriptions  as  this  aims 
to  be  may  have  three  values.  First,  they  can  present  to  all  who 
have  interest  in  such  matters  a  series  of  pictures,  as  vivid  and  real- 
istic as  possible,  of  the  vegetation  of  special  places,  and  they  are 
the  more  valuable  according  as  they  are  the  more  clearly  and  attract- 
ively written  and  the  more  aptly  illustrated.  Second,  they  should 
help  to  supply  information,  badly  needed  by  all  of  our  manuals, 
about  the  natural  habitats  of  the  lommon  or  important  species 
of  plants.  Third,  they  can  form  storehouses  of  facts  about  vege- 
tation upon  which  the  future  student  can  draw  as  the  advance  of 
physiological  ecology  gradually  makes  possible  an  understanding 
of  the  principles  underlying  physiognomic  ecology.  Such  descrip- 
tive work  can  be  done  to  profit  by  the  studt.it  whose  work  is  perforce 
confined  to  his  summer  vacations,  if  he  but  bring  to  it  time  and 
care  enough;  but  he  should  be  content  to  describe  well  and  to  leave 
interpretation  to  the  field  physiologist  yet  to  come.  Speculation 
cannot  of  itself  advance  knowledge,  and  it  can  bring  a  subject  into 
disrepute.    It  is  only,  I  believe,  through  field  physiology,  the  study 


left 


BOTANICAL  GAZETTE 


[AUGUST 


M 


in  field  laboratories  of  fundanjcntal  plar  '  namits,  that  ecological 
knowledge  can  really  be  advanced.  And  the  dynamical  problems, 
as  I  see  them,  fall  under  these  heads,  in  the  ordT  of  importance: 
(a)  physiological  life-histories  of  species,  (b)  physics  and  chemistry 
of  the  soil,  (c)  nature  of  plant  competition,  (d)  a  better  correlation 
of  meteorological  data  with  ])hysioiogical  phenomena. 

Surra  Coliege, 
Northampton,   Mass. 


